Penghusuchus

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Penghusuchus
Temporal range:
Ma[1]
Skull
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Archosauromorpha
Clade: Archosauriformes
Order: Crocodilia
Family: Gavialidae
Genus: Penghusuchus
Shan et al., 2009
Type species
Penghusuchus pani
Shan et al., 2009

Penghusuchus is an

Upper Miocene rocks of Penghu Island, off Taiwan. The taxon was described in 2009 by Shan and colleagues; the type species is P. pani.[2] It may be related to two other fossil Asian gavialids: Toyotamaphimeia machikanensis of Japan and Hanyusuchus sinensis of South China.[3] It was a medium-sized gavialid with an estimated total length of 4.5 metres (15 ft).[4]

Discovery

On 25 March 2006, on the coast of Neian, Shiyu, Penghu Islands, an excavator driver Mr. Ming-Kuo Pan found a fossilized crocodylian tooth exposed in the sandstone interlayer between basaltic rocks and then dug up a whole skeleton. The skeleton is 70% complete and was found in the Yuwentao Formation of the middle Miocene (more than 10 million years ago), and its sedimentary rocks were dated as 17-15 million years ago, according to the pollen dating in the stratum, making it one of the oldest and most complete vertebrate fossils known in Taiwan. The genus name is derived from its discovery site in Penghu, and the species name honored its discoverer, Mr. Ming-Kuo Pan. It is now considered to represent a unique and extinct gavialid clade in East Asia, along with the Pleistocene Toyotamaphimeia from Japan and Taiwan and the Holocene Hanyusuchus from South China.[3][5][6][7][8]

Morphological Description

Penghusuchus has several diagnostic characters, including: anterior process of

nasopharyngeal duct form Y-shaped ridge-like prominence on ventral surface of pterygoid; presence of five maxillary teeth within the range of the suborbital fenestra; 7th maxillary tooth is the largest in the first wave of maxillary teeth and maxilla is bulges; angular with a mid-dorsal process excluding surangular from posterodorsal border of external mandibular fenestra. Among these characters, the largest 7th maxillary tooth is only present on Pleistocene Toyotamaphimeia from Japan and Taiwan and the Holocene Hanyusuchus from South China, suggest a unique shared-featured of these three East-Asian taxa. Still, the size of Penghusuchus (4.5–5 m) is estimated smaller than Hanyusuchus and theToyotamaphimeia (may over 6 m), as well as some of the characters is differ from the latter two.[3][5][6][7][8]

Although been long classified as

exoccipital ventral process; the cranio-quadrate passage on the occipital surface is obscured by the convex process of the exoccipital; the splenial symphysis of the mandible extends to the length of about 5-7 teeth and forms a broad or narrow V in dorsal view. These characters are usually observed in gavialid, suggesting that these two East Asian taxa share mosaic features of both tomistomine and gavialid, filling the evolutionary gap of the two longeirostrine crocodylians.[8]

Based the vertebrae length, the total length of Penghusuchus is estimated as 4.5 metres. The holotype of Penghusuchus is an osteological mature individual and reached sexual maturity based on its neurocentral suture in precaudal vertebrae is closed.[4]

Phylogeny

Below is a cladogram based morphological studies comparing skeletal features that shows Penghusuchus as a member of Tomistominae, related to the false gharial:[9]

Crocodylidae

Crocodylinae

Tomistominae

Xaymacachampsa

Megadontosuchus

Kentisuchus

Maroccosuchus

Dollosuchoides

Thecachampsa

Penghusuchus

Toyotamaphimeia

Tomistoma cairense

Maomingosuchus

Tomistoma schlegelii False gharial

Gavialosuchus

Tomistoma lusitanicum

Paratomistoma

Tomistoma coppensi

Based on morphological studies of

gavialoids.[10] However, recent molecular studies using DNA sequencing have consistently indicated that the false gharial (Tomistoma) (and by inference other related extinct forms in Tomistominae) actually belong to Gavialoidea (and Gavialidae).[11][12][13][14][15][16][17]

Below is a

gavialid, related to both the gharial and the false gharial:[16]

Gavialidae

Gavialis gangeticus Gharial

Gavialis bengawanicus

Gavialis browni

Gryposuchus colombianus

Ikanogavialis

Gryposuchus pachakamue

Piscogavialis

Harpacochampsa

Toyotamaphimeia

Penghusuchus

Gavialosuchus

Tomistoma lusitanicum

Tomistoma schlegelii False gharial

Iijima and his colleagues named Hanyusuchus from Holocene South China. The phylogenetic analysis Penghusuchus pani, Hanyusuchus sinensis and Toyotamaphimeia machikanensis formed a monophyletic group.[3]

Crocodyloidea

Gavialoidea

Maroccosuchus zennaroi

Megadontosuchus arduini

Dollosuchoides densmorei

Xaymacachampsa kugleri

Kentisuchus astrei

Kentisuchus spenceri

Gavialidae

Tomistoma cairense

Tomistoma coppensi

Maomingosuchus petrolica

Tomistominae

Tomistoma schlegelii, false gharial

Tomistoma lusitanicum

Gavialosuchus eggenburgensis

Melitosaurus champsoides

Tomistoma calaritanum

Tomistoma gaudense

Thecachampsa carolinensis

Thecachampsa antiqua

Gavialinae

Paratomistoma courti

Penghusuchus pani

Toyotamaphimeia machikanensis

Hanyusuchus sinensis

Eosuchus lerichei

Eosuchus minor

Ocepesuchus eoafricanus

Eothoracosaurus mississippiensis

Thoracosaurus macrorhynchus

Thoracosaurus neocesariensis

Aktiogavialis puertoricensis

Eogavialis africanum

Argochampsa krebsi

Piscogavialis jugaliperforatus

Ikanogavialis gameroi

Siquisiquesuchus venezuelensis

Dadagavialis gunai

Gryposuchus neogaeus

Gryposuchus croizati

Aktiogavialis caribesi

Gryposuchus pachakamue

Gryposuchus colombianus

Rhamphosuchus crassidens

Myanmar gavialid

Gavialis lewisi

Gavialis bengawanicus

Gavialis gangeticus, gharial

Siwalik Gavialis

References

  1. PMID 34567843
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  2. .
  3. ^ .
  4. ^ .
  5. ^ a b Yoshitsugu, Kobayashi; Yukimitsu, Tomida; Tadao, Kamei; Taro, Eguchi (2006). "ANATOMY OF A JAPANESE TOMISTOMINE CROCODYLIAN, TOYOTAMAPHIMEIA MACHIKANENSIS (KAMEI ET MATSUMOTO, 1965), FROM THE MIDDLE PLEISTOCENE OF OSAKA PREFECTURE : THE REASSESSMENT OF ITS PHYLOGENETIC STATUS WITHIN CROCODYLIA". National Science Museum monographs. 35: i–121.
  6. ^ .
  7. ^ .
  8. ^ .
  9. .
  10. ^ Brochu, C.A.; Gingerich, P.D. (2000). "New tomistomine crocodylian from the Middle Eocene (Bartonian) of Wadi Hitan, Fayum Province, Egypt". University of Michigan Contributions from the Museum of Paleontology. 30 (10): 251–268.
  11. PMID 12775527
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  12. .
  13. .
  14. .
  15. .
  16. ^ .
  17. .